Experimental confirmation of aggressive mimicry by a coral reef fish

A number of potential mimetic relationships between coral reef fishes have been described, but the underlying mechanisms are poorly understood. Similarities in colour between species have often been attributed to aggressive mimicry (where predators resemble models in order to deceive prey), however this has not been tested. The fang blenny, Plagiotremus rhinorhynchos is a specialized predator that feeds on tissues of other fishes. Some individuals appear to mimic the harmless cleaner wrasse Labroides dimidiatus in order to deceive fish visiting cleaning stations, thereby increasing access to food. In this study, the ecological relationship between the mimic and model was examined at Kimbe Bay (Papua New Guinea) and the hypothesis that colour similarities represent facultative aggressive mimicry was experimentally evaluated. Some juveniles exhibited a striking resemblance to the juvenile colouration of the cleaner wrasse, but only when in close proximity to the wrasse and only when similar in size. As predicted for mimics, P. rhinorhynchos co-occurred with L. dimidiatus, but was rare relative to the model. Among site comparisons showed that the abundance of mimetic type blennies was positively correlated with the abundance of juvenile cleaner wrasses. Approximately 50% of all P. rhinorhynchos were found 1 m from the nearest L. dimidiatus, a distance significantly shorter than expected if they were not associated. A cleaner wrasse removal experiment was carried out to test whether the colour displayed by the blenny and its foraging success were contingent upon the presence of a model. In all cases, removal of the model prompted a rapid colour change to a general non-mimetic colouration in P. rhinorhynchos. Removal of L. dimidiatus also resulted in a ~20% reduction in the average foraging success of the blenny compared to controls, supporting the hypothesis that the blenny is a facultative aggressive mimic of the cleaner wrasse.     

Facultative mimicry: cues for colour change and colour accuracy in a coral reef fish

Mimetic species evolve colours and body patterns to closely resemble poisonous species and thus avoid predation (Batesian mimicry), or resemble beneficial or harmless species in order to approach and attack prey (aggressive mimicry). Facultative mimicry, the ability to switch between mimic and non-mimic colours at will, is uncommon in the animal kingdom, but has been shown in a cephalopod, and recently in a marine fish, the bluestriped fangblenny Plagiotremus rhinorhynchos, an aggressive mimic of the juvenile cleaner fish Labroides dimidiatus. Here we demonstrate for the first time that fangblennies adopted mimic colours in the presence of juvenile cleaner fish; however, this only occurred in smaller individuals. Field data indicated that when juvenile cleaner fish were abundant, the proportion of mimic to non-mimic fangblennies was greater, suggesting that fangblennies adopt their mimic disguise depending on the availability of cleaner fish. Finally, measurements of spectral reflectance suggest that not only do mimic fangblennies accurately resemble the colour of their cleaner fish models but also mimic other species of fish that they associate with. This study provides insights into the cues that control this remarkable facultative mimicry system and qualitatively measures its accuracy.     

Predation pressure on an imperfect Batesian micicry complex in the presence of alternative prey

Summary Differential predation pressure and the probability of predation on a Batesian mimicry complex and on alternative prey were estimatedin a field experiment. The mimicry complex was composed of a noxious model (Eleodes obscura (Say)) and a palatable mimic (Stenomorpha marginata (LeConte)). House crickets (Acheta domesticus) (Linn.) were used as alternative prey. The experiment was conducted for 23 nights in August and September to approximate the peak seasonal activity time period during which both models and mimics normally are exposed to predation while foraging and depositing eggs. Each night thirty prey in ratios of 16 models: 7 mimics: 7 crickets were exposed for 2.5 h to a suite of predators consisting of pallid bats (Antrozous pallidus), striped skunks (Mephitis mephitis) and ringtails (Bassariscus astutus) that had free access to the prey. The model-mimic ratio was similar to that found in nature. Predators obtained prey on 11 of the 23 nights and preferred the alternative prey (crickets) in proportions higher than was expected from a predation rate that was equal on all species of prey. Mimics were taken by predators at a rate proportional to their abundance, while models were taken at a rate considerably lower than their relative abundance. This suggests that at least some of the predators could distinguish between models and mimics and were willing to eat the mimics at higher frequencies than they were willing to eat the models. However, although the mimicry is not perfect with respect to the entire predator suite, the mimics still gain an advantage by resembling the models, compared to the predation levels on the alternate prey.     

Intensive cannibalism and feeding on bregmacerotids in <Emphasis Type="Italic">Champsodon snyderi</Emphasis> (Champsodontidae): evidence for pelagic predation

Analysis of the stomach contents of 1002 specimens of Champsodon snyderi (Champsodontidae) (17.3–91.2mm SL) from Tosa Bay, southern Japan, showed that species to primarily feed on crustaceans and fishes (87.9% by frequency, 37.6% by weight for the former; 17.3% and 60.7% for the latter, respectively), although fishes occurred more often in stomachs of individuals larger than 50mm SL. Champsodon snyderi ingested large prey fishes (60.5–101.0% of predator SL), the maximum weight recorded for a single ingested specimen being 50.9% of the predator weight. Mesopelagic Bregmaceros nectabanus were by far the dominant prey fish, followed by C. snyderi (cannibalism), indicating that C. snyderi leaves the bottom to feed in the pelagic environment during the night.     

Colour pattern as a single trait driving speciation in Hypoplectrus coral reef fishes?

Theory shows that speciation in the presence of gene flow occurs only under narrow conditions. One of the most favourable scenarios for speciation with gene flow is established when a single trait is both under disruptive natural selection and used to cue assortative mating. Here, we demonstrate the potential for a single trait, colour pattern, to drive incipient speciation in the genus Hypoplectrus (Serranidae), coral reef fishes known for their striking colour polymorphism. We provide data demonstrating that sympatric Hypoplectrus colour morphs mate assortatively and are genetically distinct. Furthermore, we identify ecological conditions conducive to disruptive selection on colour pattern by presenting behavioural evidence of aggressive mimicry, whereby predatory Hypoplectrus colour morphs mimic the colour patterns of non-predatory reef fish species to increase their success approaching and attacking prey. We propose that colour-based assortative mating, combined with disruptive selection on colour pattern, is driving speciation in Hypoplectrus coral reef fishes.     

Underwater observations of piranhas in western Brazil

Synopsis The behaviour of three piranha species,Serrasalmus marginatus, S. spilopleura, andPygocentrus nattereri, and their prey fishes was studied underwater in the Pantanal region, Mato Grosso, Brazil. General habits, predatory tactics, feeding behaviour, and social interactions while foraging, as well as defensive tactics of prey fishes were observed.S. marginatus is solitary whereas the other two species live in shoals; their agonistic behaviour varies accordingly, the simplest being displayed by the solitary species. Predatory tactics and feeding behaviour also vary:S. spilopleura shows the most varied diet and highly opportunistic feeding strategy, which includes aggressive mimicry. The solitaryS. marginatus, besides fin and scale-eating, occasionally cleans larger individuals ofP. nattereri. Several cichlid species display defensive tactics clearly related to piranha attacks: tail protecting, watching, and confronting the predator are the most commonly observed behaviours. Piranhas seem to strongly influence use of habitat, social structure, and foraging mode of the fish communities.     

Aggressive Mimicry in the Intra-populational Color Variation of the Tanganyikan Scale-eater Perissodus microlepis (Cichlidae)

A color morph of the Tanganyikan scale-eater, Perissodus microlepis (Cichlidae), considered to be an example of aggressive mimicry, is reported from the rocky littoral area at the southern end of Lake Tanganyika, Zambia. The anal fin of the morph is yellow, instead of the usual dark or transparent, being closely similar to that of the gregarious cyprichrominine cichlids, Cyprichromis leptosoma species complex and Paracyprichromis nigripinnis, abundant in the area. The color morph of Perissodus microlepis was observed to mingle with and attack shoals of the prey. About 24% of P. microlepis observed in less than 15m depth at Kasenga Point were the yellow-fin type. The color variation, which is likely associated with a variation in feeding mode within this population, probably evolved and has been maintained by the local community structure, which included abundant cyprichrominine fishes.     

Distance-dependent costs and benefits of aggressive mimicry in a cleaning symbiosis

In aggressive mimicry, a 'predatory' species resembles a model that is harmless or beneficial to a third species, the 'dupe'. We tested critical predictions of Batesian mimicry models, i.e. that benefits of mimicry to mimics and costs of mimicry to models should be experienced only when model and mimic co-occur, in an aggressive mimicry system involving juvenile bluestreaked cleaner wrasse (Labroides dimidiatus) as models and bluestriped fangblennies (Plagiotremus rhinorhynchos) as mimics. Cleanerfish mimics encountered nearly twice as many potential victims and had higher striking rates when in proximity to than when away from the model. Conversely, in the presence of mimics, juvenile cleaner wrasses were visited by fewer clients and spent significantly less time foraging. The benefits to mimic and costs to model thus depend on a close spatial association between model and mimic. Batesian mimicry theory may therefore provide a useful initial framework to understand aggressive mimicry.     

Aggressive mimicry in a coral reef fish: The prey's view

Since all forms of mimicry are based on perceptual deception, the sensory ecology of the intended receiver is of paramount importance to test the necessary precondition for mimicry to occur, that is, model‐mimic misidentification, and to gain insight in the origin and evolutionary trajectory of the signals. Here we test the potential for aggressive mimicry by a group of coral reef fishes, the color polymorphic Hypoplectrus hamlets, from the point of view of their most common prey, small epibenthic gobies and mysid shrimp. We build visual models based on the visual pigments and spatial resolution of the prey, the underwater light spectrum and color reflectances of putative models and their hamlet mimics. Our results are consistent with one mimic‐model relationship between the butter hamlet H. unicolor and its model the butterflyfish Chaetodon capistratus but do not support a second proposed mimic‐model pair between the black hamlet H. nigricans and the dusky damselfish Stegastes adustus. We discuss our results in the context of color morphs divergence in the Hypoplectrus species radiation and suggest that aggressive mimicry in H. unicolor might have originated in the context of protective (Batesian) mimicry by the hamlet from its fish predators rather than aggressive mimicry driven by its prey.     

A protective function for aggressive mimicry?

Mimicry often involves a protective element, whereby the risk of predation on mimics is reduced owing to their resemblance to unpalatable models. However, protection from predation has so far seemed unimportant in aggressive mimicry, where mimics are usually predators rather than prey. Here, we demonstrate that bluestriped fangblennies (Plagiotremus rhinorhynchos), which are aggressive mimics of juvenile bluestreak cleaner wrasse (Labroides dimidiatus), derive significant protection benefits from their resemblance to cleaner fish. Field observations revealed that mimetic fangblennies were chased by potential victims less often than individuals of a closely related, ecologically and behaviourally similar but non-mimetic species (Plagiotremus tapeinosoma). After attacks, proximity to models protected mimics from retaliation by victims, but the effect of colour similarity was less clear. Both colour resemblance and physical proximity to models thus appear to protect cleaner-fish mimics from aggression by potential and actual victims of their attacks. Our results suggest that the mimicry types observed in nature, which are usually distinguished on the basis of the benefits accrued to mimics, may in fact overlap greatly in the benefits provided.     

The cost of cleanerfish mimics to their models

In aggressive mimicry, a 'predatory' species resembles a model that is harmless or beneficial to a third species, the 'dupe'. Perhaps the most extraordinary case of aggressive mimicry occurs in Indo‐Pacific cleaning symbioses, where cleaner wrasses (the models) remove ectoparasites from larger fish clients. Several species of fangblennies mimic cleaners in behaviour and coloration. Instead of removing ectoparasites, however, fangblennies tear off fins, skin and scales from unsuspecting clients (the dupes). There is some debate over the extent to which cleanerfish mimics are really mimics because in some populations, the contribution of fish tissue to fangblenny diet is limited. In this study, I examine the impact of the resemblance between bluestriped fangblennies ( Plagiotremus rhinorhynchus ) and its putative model, the juvenile bluestreak cleaner wrasse ( Labroides dimidiatus ), on the model's cleaning activity to test the theoretical prediction that mimics should decrease the fitness of their models. I show that the presence of a bluestripe fangblenny in the vicinity of cleaner wrasses results in significantly lower client visit rates and inspection times compared to cleaners without a fangblenny nearby, and discuss why cleaner wrasses tolerate mimics near cleaning stations.     

Aggressive mimicry of the cleaner wrasse by Aspidontus taeniatus functions mainly for small blennies

The mimic blenny Aspidontus taeniatus Quoy & Gaimard is well known for its resemblance to the juvenile and adult cleaner wrasse Labroides dimidiatus (Valenciennes) in colour and shape. As various reef fishes including piscivores actively approach the cleaner wrasse to solicit cleaning by posing, two types of benefits have been suggested for this resemblance, that is, protective mimicry and aggressive mimicry. In aggressive mimicry, the mimic blenny is supposed to have considerable opportunities to bite the fin of deceived fishes when they pose, but some studies have confirmed that fin biting does not seem to be the main feeding tactic in the blenny in nature. Here, we examined the feeding tactics including fin biting by the mimic blenny in relation to its body size in a field observational survey in the coral reefs of Sesoko Island, Okinawa, Japan. The blenny was observed feeding mainly on four food items: the tentacles of Christmas tree worms, the mantle edges of boring clams, the demersal eggs in damselfishes’ nests and the fins of fishes. The feeding frequency by fin biting significantly decreased with body size, while that by egg predation significantly increased with body size of the blenny. When predating on eggs, the blenny was vigorously attacked by egg‐guarding fish, but often succeeded in raiding their nests by forming a feeding group. When feeding by fin biting, the blenny attacked prey fish without performing any cleaning. The ratio of fin biting was considerably higher in small‐sized blennies, suggesting reliance on this feeding tactic because of a difficulty in conducting a risky egg predation. Thus, our results suggest that the mimic blenny utilizes aggressive mimicry only when it is small as an alternative feeding tactic.     

Size-selective predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha): field stuides

Summary Studies of predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha) in a large, eutrophic lake showed that there was a clearly marked size threshold ( 160 mm SL) above which roach began to feed on mussels. Roach preying on various sizes of mussels selected them in proportions different from their abundance and accessibility in the habitat. The mean size of mussels ingested by roach of 220 mm and larger, which fed predominantly on Dreissena, closely followed the pattern expected for a constant ratio of mean prey size to mean predator mouth size = 0.59. To explain the size selection we applied an optimal foraging approach, based on the ability of different-sized fish to crush (cost) mussels of different sizes, and hence crushing resistance, and energy contents (benefit). We found that fish smaller than 160 mm, which showed no inclination to eat Dreissena, would only be able to take small mussels with a very high cost/benefit ratio. The real switch to Dreissena would be expected in fish of 230–240 mm that could take most of their prey from highly profitable, numerous, and easily accessible size classes while keeping the mean prey size at the optimal level relative to mean predator mouth size.     

Convergence of a cryptic saddle pattern in benthic freshwater fishes

Synopsis Many North American stream fishes have a similar color pattern of four dark saddles against a light background. An interesting feature of the pattern, in addition to its widespread taxonomic distribution, is its consistent configuration. The interval between the first and second saddle is usually the largest, and the last (third) interval is the smallest. All saddled North American freshwater fishes live on uneven, rocky substrates, and nearly all live in flowing water. It is hypothesized that these fishes achieve crypsis through disruptive coloration; the light spaces between the saddles mimic rocks and the dark saddles appear as shadows or gaps between rocks. Saddles are spaced unevenly because rocks in streams are a mixture of sizes; a fish that mimics a series of rocks of similar sizes is more conspicuous than one that mimics rocks of different sizes. The placement of saddles was measured on five North American species. In four of five North American species measured (a sculpin and three darters), the longest spaces are towards the head where the body is also the widest, this is thought to enhance crypsis because pieces of gravel tend to be round or square. In the madtom, the saddle pattern tends more towards even spacing. The madtom may not rely on camouflage to the same extent as other species examined because of decreased predation pressure associated with being nocturnal and possessing sharp spines and venom glands.     

Maintenance of female mimicry as a reproductive strategy in bluegill sunfish (Lepomis macrochirus)

Synopsis In species where male reproductive success is dependent on male competition and aggression, alternative reproductive patterns, thought to represent a reduction in male reproductive effort, sometimes occur. Female mimicry in bluegill sunfish (Lepomis macrochirus) is an example of an obligate alternative male strategy. Female mimics are small, sexually mature males which mimic the details of female behavior, and gain access to functional females attracted to the nests of large, aggressive territorial males. The costs of female mimicry relative to nesting male behavior are discussed and two hypotheses, deception and mutual gain, are presented to explain the tolerance and courtship of female mimics by nesting males.     

Similar yet different: differential response of a praying mantis to ant‐mimicking spiders

Batesian mimics typically dupe visual predators by resembling noxious or deadly model species. Ants are unpalatable and dangerous to many arthropod taxa, and are popular invertebrate models in mimicry studies. Ant mimicry by spiders, especially jumping spiders, has been studied and researchers have examined whether visual predators can distinguish between the ant model, spider mimic and spider non‐mimics. Tropical habitats harbour a diverse community of ants, their mimics and predators. In one such tripartite mimicry system, we investigated the response of an invertebrate visual predator, the ant‐mimicking praying mantis (Euantissa pulchra), to two related ant‐mimicking spider prey of the genus Myrmarachne, each closely mimicking its model ant species. We found that weaver ants (Oecophylla smaragdina) were much more aggressive than carpenter ants (Camponotus sericeus) towards the mantis. Additionally, mantids exhibited the same aversive response towards ants and their mimics. More importantly, mantids approached carpenter ant‐mimicking spiders significantly more than often that they approached weaver ant‐mimicking spiders. Thus, in this study, we show that an invertebrate predator, the praying mantis, can indeed discriminate between two closely related mimetic prey. The exact mechanism of the discrimination remains to be tested, but it is likely to depend on the level of mimetic accuracy by the spiders and on the aggressiveness of the ant model organism.     

Reexamination on the aggressive mimicry of the cleaner wrasseLabroides dimidiatus by the blennyAspidontus taeniatus (Pisces; Perciformes)

Field observations on feeding and related behavior of the mimic blennyAspidontus taeniatus and 3 species closely related to it, and the cleaner fish (model)Labroides dimidiatus were made at the coral reef of Sesoko Island, Okinawa, Japan, along with analysis of gut contents. The mimic blenny fed mostly on demersal eggs of fishes and tentacles of polychaetes, but it rarely tore pieces from the fins of host fishes even when they were posing for cleaning. The feeding habits of the mimic blenny are compared with those in other localities and with those of related species. It is concluded that the mimicry can hardly be regarded as an aggressive one: posing by host fishes seems to be a secondary result of the resemblance which may have developed because of the benefit for immunity from predation, and the resemblance itself prevents the blenny from becoming a specialized fin-eater because it can be easily recognized by host fishes.     

Juvenile bolas spiders attract psychodid flies

Large immature and mature female bolas spiders of the genus Mastophora attract certain male moths by aggressive chemical mimicry of those moth species' sex pheromones. These older spiders capture moths by swinging a bolas (i.e., a sticky globule suspended on a thread) at the approaching male moths. Juvenile bolas spiders do not use a bolas, but instead use their first two pairs of legs to grab prey, which our field observations suggested were primarily nematocerous Diptera. Our field experiments over a 2-year period demonstrated that juvenile bolas spiders attract moth flies (Psychodidae), with each species apparently specializing on a particular prey species. In three experiments, sticky traps containing young Mastophora phrynosoma spiderlings consistently captured significantly more male Psychoda phalaenoides than were captured on traps containing spiderlings of other Mastophora species or no spiderlings (control traps). Results from two of the three experiments suggested that Mastophora hutchinsoni spiderlings attract male Psychoda trinodulosa. Only two of our experiments included Mastophora bisaccata and those produced contrasting results. In the first experiment, M. bisaccata appeared to attract P. phalaenoides, albeit in lower numbers than were captured on traps containing M. phrynosoma. However, in a second experiment the following year, M. bisaccata spiderlings attracted Psychoda satchelli, a species that had not been caught on any traps the previous year. As suggested by a systematist four decades ago, the taxon currently called M. bisaccata may consist of two or more sibling species, which could account for the contrasting results obtained from our two experiments involving M. bisaccata. This is the first reported evidence that, during early developmental stadia before these spiders attract moths, juvenile bolas spiders attract their prey.     

Ultraviolet Radiation Absorbance by Coral Reef Fish Mucus: photo-protection and Visual Communication

Tropical reef fishes are exposed to high levels of damaging ultraviolet radiation. Here we report the widespread distribution of both UVA- and UVB-absorbing compounds in the epithelial mucus of these fishes. Mucus from 137 reef fish species was examined by spectrophotometry and 90% were found to have strong absorbance peaks between 290 and 400nm. Most fish species (78%) had more than one peak, that suggests a broad-band ultraviolet screening function for their mucus. Thalassoma duperrey, a tropical wrasse, was able to alter the absorbance of its epithelial mucus in response to both naturally and experimentally manipulated UV regimes. Visual modeling suggests that a fish with UV vision, such as Dascyllus albisella, could detect the changes in mucus spectra of T. duperrey that occurred in these experiments.     

Observations on the distribution patterns,behaviour, diets and reproductive cycles of sand-dwelling clinids (Perciformes: Clinidae) from South Africa

Synopsis Twenty-two samples of sand-dwelling fishes were collected from sublittoral sandy substrata off the Cape Peninsula, South Africa using a novel quantitative technique in which the ichthyocide, rotenone, was introduced beneath 6.25 m² plastic sheets weighted around the perimeter with chain. A total of 94 fish of eight species and four families were recorded. Of these, four species of the family Clinidae made up 92% of the material. Fish densities varied from 0.10–2.96 fish m^–2 (0.17–1.14 g m^–2). Distribution patterns amongst the Clinidae were explained by sediment particle size, with all of the newly discovered species, Cancelloxus longior, inhabiting fine sand (median grain size 0.25–0.50 mm), and all Xenopoclinus leprosus, C. elongatus and Pavoclinus smalei occurring in gravel (> 1 mm). Xenopoclinus kochi was found in all sediment types, but was most abundant in coarse substrata. Diets of all four species were similar, consisting chiefly of amphipods and isopods. However, small differences in prey preference were evident. All four species reached sexual maturity at 25–30 mm standard length, were viviparous, exhibited superembryonation and gave birth to live young of approximately 14 mm. Breeding seasonality occurred in all species except P. smalei.